Ennaltamäärätty kääntymys: Predestinaatio ja vapaan tahdon ongelma Jonathan Edwardsin teologiassa

Tämän tutkielman tehtävänä on ollut selvittää onko ihmisellä vapaata tahtoa Jonathan Edwardsin mukaan. Lähteenäni minulla on ollut Edwardsin Freedom of the Will -teos sekä Miscellaneous- sarjasta Concerning the Divine Degrees -kirjoitus. Metodina minulla on ollut systemaattinen analyysi. Edwardsin taustana oli puritanismi. Esimerkiksi sakramenteilla ei ollut puritanismin piirissä samaa ratkaisevaa merkitystä uskon synnyttäjänä ja ylläpitäjänä kuin perinteisissä kristillisissä kirkoissa oli ollut. Puritanistien keskeinen armonväline oli saarna. Saarnalla pyrittiin vaikuttamaan ihmisen tahtoon antamalla rationaalisia perusteita Raamatun pelastustapahtumasta. Tämä on nähtävissä myös Jonathan Edwardsin teologiassa. Deismi ja valistus vaikuttivat myös Edwardsin aikaiseen ajatteluun voimakkaasti. Edwardsin Berkeleytä muistuttava idealismi kuitenkin takasi sen, että Jumala ei irronnut maailmasta vaan oli kaikessa koko ajan läsnä. Erityisesti Locke Newtonin luonnontieteiden filosofina vaikutti Edwardsin filosofiseen ajatteluun tahdon vapaus -kysymyksen osalta. Edwards pyrki Locken ja Newtonin avulla puolustamaan perinteistä kalvinistista kristillisyyttä. Tämän kilpailijaksi puritanismin piirissä oli tullut ajan henkeen sopiva järkiperäinen arminiolaisuus, joka edusti myös anti-revivalismia. Tähän arminiolaisuuteen liittyi perinteisistä kristillisistä opeista luopuminen. Tämä arminiolaisuus oli kehittynyt antropologisempaan suuntaan Remonstranssin ajoista korostaen ihmisen vapaita tahdon akteja. Näin ihminen pystyi päättämään uskoiko hän vai ei. Edwardsille usko ja pelastuminen perustuivat Jumalan valintaan, jossa kolmiyhteinen Jumala otti ihmisen yhteyteensä. Tämä kääntymys ilmeni kommuuniona ja uniona Kristuksen kanssa. Edwardsin ajattelua tahdon vapauden osalta avaa kompatibilismi. Kompatibilismin mukaan ennaltamäärääminen ei ole ristiriidassa vapaan tahdon kanssa, vaan päinvastoin sen lähtökohta ja perusta. Kääntymissä olemme eri näkökulmista täysin passivisia ja täysin aktiivisia. Kompatibilismin myötä Edwardsin kirjoitusten pohjalta pystytään johtamaan ennalta määrätty uskonratkaisu. Myöhemmin Edwardsin seuraajat New Lights harppasivat ironisesti arminiolaiseen suuntaan korostaen juuri uskonratkaisua. Vaikka Jumalalla on sanan varsinaisessa merkityksessä vapaa tahto, niin silti Hän toimii välttämättä pyhästi. Samoin Jeesus Kristus on linkkinä ihmisten välttämättömälle, mutta vastuulliselle toiminnalle. Jeesuksen valinta on kaikkien pyhien valinnan perusta. Ne, jotka Jumala hylkää helvettiin saavat oikeudenmukaisen rangaistuksen, koska Jumala rankaisee heissä itsessään olevaa pahaa. Se, miksi Jumala valitsee toiset ja hylkää toiset, kuuluu Jumalan salattuun tahtoon. Jumalan ilmoitetun tahdon mukaan Hän tahtoo kaikkien pelastuvan. Arminiolaiset hylkäävät ennaltamääräämisen, mutta hyväksyvät ennalta tietämisen. Edwardsille nämä ovat sama asia. Edwardsin mukaan välttämättömyys ei ole ristiriidassa vapauden kanssa. Luonnollisella ja moraalisella välttämättömyydellä on yhteys. Arkikielessä moraalista välttämättömyyttä ei oleteta. Koko tahdon vapaus -kysymys on pitkälti kielellinen. Arkikielessä ja filosofisessa kielessä on ero. Tahto ei ole agentti, joka voi valita itsenäisesti. Vapaus on ihmisen, ei tahdon ominaisuus. Siksi kysymys tahdon vapaudesta tulisi esittää onko ihminen vapaa. Edwards ja arminiolaiset ymmärtävät vapauden eri tavoin. Edwardsin mukaan tahdolla on oltava alku, eikä tahto voi koskaan arminiolaisten tavoin neutraali. Arminiolaiset hylkäävät perisyntiopin klassisessa merkityksessä, mistä seuraa erilainen lähtökohta. Sellaisena kuin arminiolaiset esittävät vapaan tahdon, meillä ei sitä Edwardsin mukaan ole. Vaikka kaikki tapahtuu Edwardsin mukaan välttämättä, niin silti olemme vastuullisia moraalisia agentteja.

Reinhold Niebuhrin syntikäsitys

Reinhold Niebuhrin syntikäsitys rakentuu hänen ihmiskäsitykseensä kuuluvan dikotomian henki-luonto, sekä sitä sisällöllisesti vastaavan käsiteparin vapaus-rajallisuus, varaan. Ihmisen hengen kykyjä ovat transsendenssi yli luonnon, itsetranssendenssi sekä vapaus. Toisaalta ihminen on luontonsa kautta sidottu elollisen luonnon rajallisuuteen, vajavaisuuteen ja kuolevaisuuteen. Ihmisen rajallisuus on hengen kykyjen tavoin Jumalan luomisessa ihmiselle antama ominaisuus, ja siksi hyvä. Ihmisen asema rajallisena ja vapaana, eli ihmisen ristiriita antaa hänelle aiheen synnin tekemiseen. Tämä ihmisen ristiriita ei kuitenkaan ole synnin syy. Ihmisen asema luonnon ja hengen rajapinnassa saa hänet levottomaksi. Levottomuus on ihmisen ristiriidan väistämätön seuralainen ja samalla synnin sisäinen ennakkoehto. Raamatun lankeemuskertomus on Niebuhrin mukaan myytti, joka kuvaa symbolisesti ihmisen lankeamista syntiin. Lankeemusmyyttiä ei pidä tulkita kirjaimellisesti eikä kausaalisesti, sillä tällöin se menettää todellisen merkityksensä. Lankeemusmyytin mukaan ihmisen pahaa edeltää pahan voima, eli paholainen. Koska paholainen lankesi ennen ihmistä, ihmisen synti ei seuraa väistämättä ihmisen tilanteesta vapaana ja rajallisena. Ihmisen synti ei myöskään ole silkkaa perverssiyttä, tietoista pahan tekemistä ja Jumalan vastaista uhmaa. Ihmisen vapaus ja rajallisuus muuttuvat kiusaukseksi vasta, kun ne on tulkittu väärin. Ihmisen tekoa edeltävä pahan voima ehdottaa tätä tulkintaa ihmiselle. Niebuhr kuvaa Kierkegaardilta lainaamillaan ilmauksilla, kuinka 'synti asettaa itsensä' ja 'synti edellyttää itsensä'. Hän puhuu myös 'synnin laadullisesta hypystä'. Syntiä ei koskaan voi johtaa tietystä tilanteesta nousevasta kiusauksesta. Itsensä asettava synti on ihmisen oma synti. Synti edellyttää itsensä tarkoittaa, että kaikkea syntiä edeltää epäuskon synti. Ilman epäuskoa ihmisen levottomuus ei johtaisi syntiin. Niebuhrin metodinen tukeutuminen Kierkegaardiin ei ole johdonmukaista, mistä osittain johtuu, että hänen käsityksensä ihmisen syntiin lankeamisesta on ristiriitainen. Niebuhr sotkee myytti- ja historiapuheen keskenään. Ihmisen tietoisuus ja vapaus ovat edellytys sille, että ihminen ylipäänsä voi tehdä syntiä. Niebuhr torjuu perinteisen kristillisen näkemyksen, jonka mukaan lankeemus merkitsi radikaalia muutosta ihmisen tahdon vapauteen. Ihmisen huomio, ettei hän kykene tekemään valintaa hyvän ja pahan välillä, on Niebuhrin mukaan merkittävin osoitus ihmisen vapaudesta. Ihmisen tietoisuus synnistään ei koskaan ole täydellistä. Ihminen ei tee syntiä täysin tietoisesti ja harkitusti. Toisaalta ihmisen osittainen tietämättömyys synnistään on synnin seuraus. Voidakseen jatkaa synnin tekemistä, eli voidakseen pitäytyä kohtuuttomassa itserakkaudessaan, ihmisen on petettävä ensin itseään, sitten toisia ihmisiä. Niebuhr määrittelee hyvin pidättyväisesti näkemyksensä perisynnistä. Perisynti on ennen kaikkea ihmisen taipumus synnin tekemiseen. Synti on peräisin ihmisen tahdon viasta. Toisaalta Niebuhrin mukaan ihminen tekee väistämättä syntiä. Niebuhr on muutamissa yhteyksissä varovaisesti sillä kannalla, että ihmisen luonto on synnin seurauksena turmeltunut. Toisissa yhteyksissä hän torjuu tällaisen näkemyksen. Niebuhrin näkemys synnin väistämättömyydestä sekä näkemys ihmisen tahdon viasta jäävät jossain määrin irrallisiksi osiksi hänen ihmiskäsityksessään. Ihmisen synti on ylpeyttä ja aistillisuutta. Ihmisellä on taipumus kieltää joko rajallisuutensa tai vapautensa. Kieltäessään rajallisuutensa hän lankeaa ylpeyteen. Kieltäessään vapautensa hän lankeaa aistillisuuteen. Ylpeyden muotoja on neljä: vallanylpeys, tiedonylpeys, moraalinen ylpeys ja hengellinen ylpeys. Aistillisuudessa on kolme vaihetta. Aistillisuus on itserakkautta, yritystä paeta itseä toiseen ihmiseen tai luontoon sekä kolmanneksi pakoa tyhjyyteen ja tiedostamattomuuteen. Synti ja syyllisyys liittyvät väistämättä seksiin. Seksi on aistillisuuden elävin ilmaus ja samalla ihmisen luovuuden kliimaksi. Niebuhr pitää ihmisyhteisöjä yksilöiden ohella rajallisessa mielessä synnin subjekteina. Yhteisöillä on yksilöiden tavoin henki ja luonto, sekä itsetranssendenssin elimiä. Niebuhr pitää yhteisöjä yksilöitä moraalisesti pahempina. Yhteisöt aiheuttavat yksilöitä enemmän objektiivista pahaa. Niebuhr kritisoi syntikäsityksensä pohjalta liberalismiksi kutsumaansa ajattelua. Liberalismin perustana on Niebuhrin mukaan kehityksen idea, joka on perusteetonta optimistista uskoa täydellistyvään maailmaan ja uskoa historiaan itsensä lunastavana prosessina. Niebuhrin mukaan kaikki vallankäyttö elämässä johtaa sortoon ja epäoikeudenmukaisuuteen. Sen vuoksi yhteisöjä on hallittava voimien balanssilla sekä hallituksella. Nämä tekijät on mahdollista ottaa pätevästi huomioon vain demokratiassa. Niebuhr näkee kirkon silkkana ihmisyhteisönä, jolle on kuitenkin suotu evankeliumi. Hän korostaa kirkon syntisyyttä, muttei ollenkaan sen pyhyyttä.

Diakonian työalajohtaja ja hänen tiimiläistensä työhyvinvointi

Tutkielman aiheena on kuvata ja analysoida seurakunnan työalajohtajuutta työhyvinvoinnin näkökulmasta. Aineistona on kahdentoista diakonian työalajohtajan teemahaastattelut. Tutkimuskysymyksinä ovat: 1.Millaisilla edellytyksillä työalajohtaja voi toimia seurakunnassa? 2.Mistä työhyvinvointi diakoniatyössä haastateltavien kokemuksen mukaan muodostuu? 3.Millaisilla toimenpiteillä työalajohtajat edistävät tiimiläistensä työhyvinvointia? Sisällönanalyysilla saatujen tulosten mukaan työalajohtajuus onnistuu, kun kirkkoherra on työalajohtajuuden tukena ja vallan ja vastuun delegointi on selkeää ja riittävän laajaa. Ilman ylemmän johdon tukea työalajohtajan tehtävässä toimiminen on vaikeaa tai jopa mahdotonta. Työalajohtajan toimivalta ja vastuu tuli määritellä selkeästi. Kirkkoherralla oli merkittävä rooli myös työyhteisön suhtautumisen muokkaajana. Huomattavan monessa seurakunnassa varsinkin papeilla oli ollut vaikeuksia suhtautua johtajuuteen aseman sijasta tehtävänä. Haastatellut työalajohtajat painottivat johtajuutta ammattina, johon voi ja pitää kouluttautua sekä toisaalta prosessina, jossa ei tule valmiiksi. Työalajohtajan tuli haastateltavien mukaan olla työalansa substanssiosaaja. Diakoniatyössä monet työn piirteet toimivat sekä työhyvinvointia edistävinä että sitä haastavina. Työn haasteellisuus ja monipuolisuus tuottivat työniloa, mutta tuntuivat myös raskailta. Työn rajaamisen ongelmat olivat ajankohtaisia kaikissa haastatteluseurakunnissa. Työajattomuus koettiin pääsääntöisesti voimavaraksi ja mahdollisuudeksi. Verkostoyhteistyö vei voimavaroja, mutta antoi myös tunteen yhdessä tehtävästä laajemmasta työstä. Merkittävä vaikutus työhyvinvointiin oli työntekijän suhtautumistavalla työhönsä. Työalajohtajat edistivät tiimiläistensä työhyvinvointia luomalla rakenteita ja toimintaedellytyksiä työn kehittämiselle ja työn tekemiselle. Yhteisöllisyyttä vahvistavat toimenpiteet liittivät sooloilevat työntekijät osaksi tiimiä. Hoitamalla hallintoon liittyvät tehtävät työalajohtajat vapauttivat tiimiläisten työpanosta varsinaisen diakoniatyön tekemiseen. Työalajohtajat toimivat diakonian äänenä välittämällä tietoa tiimin ja hallinnon välillä molempiin suuntiin sekä tuomalla esiin diakonian näkökulmaa seurakunnassa ja toiminta-alueella. Työalajohtajat edistivät tiimin keskusteluyhteyttä ja huolehtivat tiimin työskentelyilmapiiristä. Keskustelujen avulla he loivat yhteisiä toimintatapoja ja linjanvetoja sekä vahvistivat maltillista suhtautumista työhön. Diakonian työalajohtajista löytyi neljä työalajohtajtyyppiä suhteessa työhyvinvoinnin edistämiseen: Koordinoija, Pelinrakentaja, Majakka ja Opas. Koordinoijat korostivat toimivien rakenteiden merkitystä. Pelinrakentajat painottivat tiimin yhteistä suunnittelua, jakamista ja arviointia. Majakka-työalajohtajat toimivat vakaina yhteisöllisyyden ja optimismin vahvistajina. Oppaat loivat levollista ja laadukasta työskentelyilmapiiriä. Diakonian työalajohtajat pyrkivät omalla johtamistoiminnallaan luomaan tiimiläisilleen mahdollisimman hyvät puitteet tehdä työtä hyvällä mielellä. Avainsanat – Nyckelord kirkon työntekijät, seurakunnat, seurakuntatyö, seurakuntahallinto, diakonia, johtajuus, esimiehet, johtaminen, organisaatiokulttuuri, työyhteisöt, työhyvinvointi, työn organisointi, työilmapiiri

Generations and turnout : The generational effect in electoral participation in Finland

The relationship between age and turnout has been curve-linear as electoral participation first increases with age, remains relatively stable throughout middle-age and then gradually declines as certain physical infirmities set in (see e.g. Milbrath 1965). Alongside this life-cycle effect in voting, recent pooled cross-sectional analyses (see e.g. Blais et al. 2004; Lyons and Alexander 2000) have shown that there is also a generational effect, referring to lasting differences in turnout between various age groups. This study firstly examines the extent to which the generational effect applies in the Finnish context. Secondly, it investigates the factors accounting for that effect. The first article, based on individual-level register data from the parliamentary elections of 1999, shows that turnout differences between the different age groups would be even larger if there were no differences in social class and education. The second article examines simultaneously the effects of age, generation and period in the Finnish parliamentary elections of 1975-2003 based on pooled data from Finnish voter barometers (N = 8,634). The results show that there is a clear life cycle, generational and period effect. The third article examines the role of political socialisation in accounting for generational differences in electoral participation. Political socialisation is defined as the learning process in which an individual adopts various values, political attitudes, and patterns of actions from his or her environment. The multivariate analysis, based on the Finnish national election study 2003 (N=1,270), indicated that if there were no differences in socialisation between the youngest and the older generations, the difference in turnout would be much larger than if only sex and socioeconomic factors are controlled for. The fourth article examines other possible factors related to generational effect in voting. The results mainly apply to the Finnish parliamentary elections of 2003 in which we have data available. The results show that the sense of duty by far accounts for the generational effect in voting. Political interest, political knowledge and non-parliamentary participation also narrowed the differences in electoral participation between the youngest and the second youngest generations. The implication of the findings is that the lower turnout among the current youth is not a passing phenomenon that will diminish with age. Considering voting a civic duty and understanding the meaning of collective action are both associated with the process of political socialisation which therefore has an important role concerning the generational effect in turnout.

Essays on macroeconomic effects of fiscal policy rules

Economic and Monetary Union can be characterised as a complicated set of legislation and institutions governing monetary and fiscal responsibilities. The measures of fiscal responsibility are to be guided by the Stability and Growth Pact, which sets rules for fiscal policy and makes a discretionary fiscal policy virtually impossible. To analyse the effects of the fiscal and monetary policy mix, we modified the New Keynesian framework to allow for supply effects of fiscal policy. We show that defining a supply-side channel for fiscal policy using an endogenous output gap changes the stabilising properties of monetary policy rules. The stability conditions are affected by fiscal policy, so that the dichotomy between active (passive) monetary policy and passive (active) fiscal policy as stabilising regimes does not hold, and it is possible to have an active monetary - active fiscal policy regime consistent with dynamical stability of the economy. We show that, if we take supply-side effects into ac-count, we get more persistent inflation and output reactions. We also show that the dichotomy does not hold for a variety of different fiscal policy rules based on government debt and budget deficit, using the tax smoothing hypothesis and formulating the tax rules as difference equations. The debt rule with active monetary policy results in indeterminacy, while the deficit rule produces a determinate solution with active monetary policy, even with active fiscal policy. The combination of fiscal requirements in a rule results in cyclical responses to shocks. The amplitude of the cycle is larger with more weight on debt than on deficit. Combining optimised monetary policy with fiscal policy rules means that, under a discretionary monetary policy, the fiscal policy regime affects the size of the inflation bias. We also show that commitment to an optimal monetary policy not only corrects the inflation bias but also increases the persistence of output reactions. With fiscal policy rules based on the deficit we can retain the tax smoothing hypothesis also in a sticky price model.

Tibetan Market Participation in China

Since the Chinese government began implementing economic reforms in the late 1970s, China has experienced profound economic change and growth. Like other parts of China, Tibetan areas of China have also experienced wide-ranging economic change with growth even higher than the China-wide average in certain years. Though China s strategic policy of developing the West provided many opportunities for economic and business activities, Tibetans have proven poorly equipped to respond to and take advantage of these opportunities. This study is about people, about market participation and specifically about why Tibetans do not effectively participate in the market in the context of China s economic development process. Many political, social, cultural and environmental factors explain the difficulties met by Tibetan communities. However, this study focuses on three factors: the social and culture context, government policy and education. The Buddhistic nature of Tibetan communities, particularly the political and economic system in traditional Tibetan society, explains this, especially after implementation of new national economic policies. An inclusive economic development policy that promotes local people s participation in the market demands serious consideration of local conditions. Unfortunately, such considerations often ignore local Tibetan realities. The economic development policy in Tibetan areas in China is nearly always an attempt to replicate the inland model and open up markets, even though economic and sociopolitical conditions in Tibet are markedly unlike much of China. A consequence of these policies is increasing numbers of non-Tibetan migrants flowing into Tibetan areas with the ensuing marginalization of Tibetans in the marketplace. Poor quality education is another factor contributing to Tibetan inability to effectively participate in the market. Vocational and business education targeting Tibetans is of very low quality and reflective of government failing to consider local circumstances when implementing education policy. The relatively few Tibetans who do receive education are nearly always unable to compete with non-Tibetan migrants in commercial activity. Encouraging and promoting Tibetan participation in business development and access to quality education are crucial for a sustainable and prosperous society in the long term. Particularly, a localized development policy that considers local environmental conditions and production as well as local culture is crucial. Tibet s economic development should be based on local environmental and production conditions, while utilizing Tibetan culture for the benefit of creating a sustainable economy. Such a localized approach best promotes Tibetan market participation. Keywords: Tibet cultural policy education market participation

Bending the Rules of Cell Protrusions : Molecular Mechanisms and Biological Roles of Inverse-BAR Proteins in Cell Morphogenesis

Plasma membrane adopts myriad of different shapes to carry out essential cellular processes such as nutrient uptake, immunological defence mechanisms and cell migration. Therefore, the details how different plasma membrane structures are made and remodelled are of the upmost importance. Bending of plasma membrane into different shapes requires substantial amount of force, which can be provided by the actin cytoskeleton, however, the molecules that regulate the interplay between the actin cytoskeleton and plasma membrane have remained elusive. Recent findings have placed new types of effectors at sites of plasma membrane remodelling, including BAR proteins, which can directly bind and deform plasma membrane into different shapes. In addition to their membrane-bending abilities, BAR proteins also harbor protein domains that intimately link them to the actin cytoskeleton. The ancient BAR domain fold has evolved into at least three structurally and functionally different sub-groups: the BAR, F-BAR and I-BAR domains. This thesis work describes the discovery and functional characterization of the Inverse-BAR domains (I-BARs). Using synthetic model membranes, we have shown that I-BAR domains bind and deform membranes into tubular structures through a binding-surface composed of positively charged amino acids. Importantly, the membrane-binding surface of I-BAR domains displays an inverse geometry to that of the BAR and F-BAR domains, and these structural differences explain why I-BAR domains induce cell protrusions whereas BAR and most F-BAR domains induce cell invaginations. In addition, our results indicate that the binding of I-BAR domains to membranes can alter the spatial organization of phosphoinositides within membranes. Intriguingly, we also found that some I-BAR domains can insert helical motifs into the membrane bilayer, which has important consequences for their membrane binding/bending functions. In mammals there are five I-BAR domain containing proteins. Cell biological studies on ABBA revealed that it is highly expressed in radial glial cells during the development of the central nervous system and plays an important role in the extension process of radial glia-like C6R cells by regulating lamellipodial dynamics through its I-BAR domain. To reveal the role of these proteins in the context of animals, we analyzed MIM knockout mice and found that MIM is required for proper renal functions in adult mice. MIM deficient mice displayed a severe urine concentration defect due to defective intercellular junctions of the kidney epithelia. Consistently, MIM localized to adherens junctions in cultured kidney epithelial cells, where it promoted actin assembly through its I-BAR andWH2 domains. In summary, this thesis describes the mechanism how I-BAR proteins deform membranes and provides information about the biological role of these proteins, which to our knowledge are the first proteins that have been shown to directly deform plasma membrane to make cell protrusions.

Net neutrality in Australia: The debate continues, but no policy in sight

The ways in which Internet traffic is managed have direct consequences on Internet users’ rights as well as on their capability to compete on a level playing field. Network neutrality mandates to treat Internet traffic in a non-discriminatory fashion in order to maximise end users’ freedom and safeguard an open Internet.

GDNF family receptors in peripheral target innervation and hormone production

Glial cell line-derived neurotrophic factor (GDNF) family ligands: GDNF, neurturin, persephin and artemin, signal through a receptor tyrosine kinase Ret by binding first to a co-receptor (GFRα1-4) that is attached to the plasma membrane. The GDNF family factors can support the survival of various peripheral and central neuronal populations and have important functions also outside the nervous system, especially in kidney development. Activating mutations in the RET gene cause tumours in neuroendocrine cells, whereas inactivating mutations in RET are found in patients with Hirschsprung s disease (HSCR) characterized by loss of ganglionic cells along the intestine. The aim of this study was to examine the in vivo functions of neurturin receptor GFRα2 and persephin receptor GFRα4 using knockout (KO) mice. Mice lacking GFRα2 grow poorly after weaning and have deficits in parasympathetic and enteric innervation. This study shows that impaired secretion of the salivary glands and exocrine pancreas contribute to growth retardation in GFRα2-KO mice. These mice have a reduced number of intrapancreatic neurons and decreased cholinergic innervation of the exocrine pancreas as well as reduced excitatory fibres in the myenteric plexus of the small intestine. This study also demonstrates that GFRα2-mediated Ret signalling is required for target innervation and maintenance of soma size of sympathetic cholinergic neurons and sensory nociceptive IB4-binding neurons. Furthermore, lack of GFRα2 in mice results in deficient perception of temperatures above and below thermoneutrality and in attenuated inflammatory pain response. GFRα4 is co-expressed with Ret predominantly in calcitonin-producing thyroid C-cells in the mouse. In this study GFRα4-deficient mice were generated. The mice show no gross developmental deficits and have a normal number of C-cells. However, young but not adult mice lacking GFRα4 have a lower production of calcitonin in thyroid tissue and consequently, an increased bone formation rate. Thus, GFRα4/Ret signalling may regulate calcitonin production. In conclusion, this study reveals that GFRα2/Ret signalling is crucial for the development and function of specific components of the peripheral nervous system and that GFRα4-mediated Ret signalling is required for controlling transmitter synthesis in thyroid C-cells.

Responses of Scots pine to nickel and copper exposure and herbivory

The goal of this thesis was to examine the ecophysiological responses of Scots pine (Pinus sylvestris L.), with an emphasis on the oxidative enzyme peroxidase and plant phenolics to environmental stresses like elevated levels of nickel (Ni) and copper (Cu), and herbivory. The effects of Ni and Cu were studied in a gradient survey at a sulphur dioxide contaminated site in the Kola Peninsula, and with experiments in which seedlings were exposed to Ni mist or to Ni and Cu amended into the soil. In addition, experimental Ni exposure was combined with disturbance of the natural lichen cover of the forest ground layer. Pine sawfly attack was simulated in the early season defoliation experiment, in which mature Scots pine were defoliated (100 %) during two successive years in a dry, nutrient-poor Scots pine stand. In addition, the effect of previous defoliation on the growth of sawfly (Diprion pini L.) larvae was studied. Apoplastic peroxidase activity was elevated in the needles of pine in a Ni- , Cu- and SO2- polluted environment, which indicated an increased oxidative stress. Increased foliar peroxidase activity due to Ni contamination was shown in the experiment, in which Ni was added as mist. No such response was found in peroxidase acitivity of the roots exposed to elevated Ni and/or Cu in the soil. Elevated Ni in the soil increased the concentration of foliar condensed tannins, which are able to bind heavy metals in the cells. Addition of low levels of Ni in the soil appeared to benefit pine seedlings, which was seen as promoted shoot growth and better condition of the roots. Wet Ni deposition of 2000 mg m-2 reduced growth and survival of pine seedlings, whereas deposition levels 200 mg m-2 or 20 mg m-2 caused no effects in a 2-y lasting experiment. The lichen mat on the forest floor did not act as an effective buffer against the adverse impacts of heavy metals on pine seedlings. However, some evidence was found indicating that soil microbes profited from the lichen mat. Artificial defoliation increased peroxidase activity in the Scots pine needles. In addition, defoliation decreased nitrogen, diamine putrescine and glucose concentrations in the needles and increased the concentrations of several phenolic compounds, starch and sucrose. Previous artificial defoliation led to poor growth of sawfly larvae reared on the pines, suggesting delayed induced resistance in Scots pine. However, there was no consistent relationship between inducibility (proportional increase in a compound following defoliation) and adverse effects on the growth of pine sawfly larvae. The observed inducible responses in needle phenolics due to previous defoliation thus appear to represent non-specific responses against sawflies.

Short- and long-term consequences of food resources on Ural owl Strix uralensis reproduction

Life-history theory states that although natural selection would favour a maximisation of both reproductive output and life-span, such a combination can not be achieved in any living organism. According to life-history theory the reason for the fact that not all traits can be maximised simultaneously is that different traits compete with each other for resources. These relationships between traits that constrain the simultaneous evolution of two or more traits are called trade-offs. Therefore, during different life-stages an individual needs to optimise its allocation of resources to life-history components such as growth, reproduction and survival. Resource limitation acts on these traits and therefore investment in one trait, e.g. reproduction, reduces the resources available for investment in another trait, e.g. residual reproduction or survival. In this thesis I study how food resources during different stages of the breeding event affect reproductive decisions in the Ural owl (Strix uralensis) and the consequences of these decisions on parents and offspring. The Ural owl is a suitable study species for such studies in natural populations since they are long-lived, site-tenacious, and feed on voles. The vole populations in Fennoscandia fluctuate in three- to four-year cycles, which create a variable food environment for the Ural owls to cope with. The thesis gives new insight in reproductive costs and their consequences in natural animal populations with emphasis on underlying physiological mechanisms. I found that supplementary fed Ural owl parents invest supplemented food resources during breeding in own self-maintenance instead of allocating those resources to offspring growth. This investment in own maintenance instead of improving current reproduction had carry-over effects to the following year in terms of increased reproductive output. Therefore, I found evidence that reduced reproductive costs improves future reproductive performance. Furthermore, I found evidence for the underlying mechanism behind this carry-over effect of supplementary food on fecundity. The supplementary-fed parents reduced their feeding investment in the offspring compared to controls, which enabled the fed female parents to invest the surplus resources in parasite resistance. Fed female parents had lower blood parasite loads than control females and this effect lasted until the following year when also reproductive output was increased. Hence, increased investment in parasite resistance when resources are plentiful has the potential to mediate positive carry-over effects on future reproduction. I further found that this carry-over effect was only present when potentials for future reproduction were good. The thesis also provides new knowledge on resource limitation on maternal effects. I found that increased resources prior to egg laying improve the condition and health of Ural owl females and enable them to allocate more resources to reproduction than control females. These additional resources are not allocated to increase the number of offspring, but instead to improve the quality of each offspring. Fed Ural owl females increased the size of their eggs and allocated more health improving immunological components into the eggs. Furthermore, the increased egg size had long-lasting effects on offspring growth, as offspring from larger eggs were heavier at fledging. Limiting resources can have different short- and long-term consequences on reproductive decisions that affect both offspring number and quality. In long-lived organisms, such as the Ural owl, it appears to be beneficial in terms of fitness to invest in long breeding life-span instead of additional investment in current reproduction. In Ural owls, females can influence the phenotypic quality of the offspring by transferring additional resources to the eggs that can have long-lasting effects on growth.

Evolutionary Genetics in the WIld : from Populations to Individuals

Predicting evolutionary outcomes and reconstructing past evolutionary transitions are among the main goals of evolutionary biology. Ultimately, understanding the mechanisms of evolutionary change will also provide answers to the timely question of whether and how organisms will adapt to changing environmental conditions. In this thesis, I have investigated the relative roles of natural selection, random genetic drift and genetic correlations in the evolution of complex traits at different levels of organisation from populations to individuals. I have shown that natural selection has been the driving force behind body shape divergence of marine and freshwater threespine stickleback (Gasterosteus aculeatus) populations, while genetic drift may have played a significant role in the more fine scale divergence among isolated freshwater populations. These results are concurrent with the patterns that have emerged in the published studies comparing the relative importance of natural selection and genetic drift as explanations for population divergence in different traits and taxa. I have also shown that body shape and armour divergence among threespine stickleback populations is likely to be biased by the patterns of genetic variation and covariation. Body shape and armour variation along the most likely direction of evolution the direction of maximum genetic variance reflects the general patterns of variation observed wild populations across the distribution range of the threespine stickleback. Conversely, it appears that genetic correlations between the sexes have not imposed significant constraints on the evolution of sexual dimorphism in threespine stickleback body shape and armour. I have demonstrated that the patterns of evolution seen in the wild can be experimentally recreated to tease out the effects of different selection agents in detail. In addition, I have shown how important it is to take into account the correlative nature of traits, when making interpretations about the effects of natural selection on individual traits. Overall, this thesis provides a demonstration of how considering the relative roles of different mechanism of evolutionary change at different levels of organisation can aid in an emergence of a comprehensive picture of how adaptive divergence in wild populations occurs.

Electron cryo-microscopy studies of bacteriophage phi8 and archaeal virus SH1

Symmetry is a key principle in viral structures, especially the protein capsid shells. However, symmetry mismatches are very common, and often correlate with dynamic functionality of biological significance. The three-dimensional structures of two isometric viruses, bacteriophage phi8 and the archaeal virus SH1 were reconstructed using electron cryo-microscopy. Two image reconstruction methods were used: the classical icosahedral method yielded high resolution models for the symmetrical parts of the structures, and a novel asymmetric in-situ reconstruction method allowed us to resolve the symmetry mismatches at the vertices of the viruses. Evidence was found that the hexameric packaging enzyme at the vertices of phi8 does not rotate relative to the capsid. The large two-fold symmetric spikes of SH1 were found not to be responsible for infectivity. Both virus structures provided insight into the evolution of viruses. Comparison of the phi8 polymerase complex capsid with those of phi6 and other dsRNA viruses suggests that the quaternary structure in dsRNA bacteriophages differs from other dsRNA viruses. SH1 is unusual because there are two major types of capsomers building up the capsid, both of which seem to be composed mainly of single beta-barrels perpendicular to the capsid surface. This indicates that the beta-barrel may be ancestral to the double beta-barrel fold.

Dynamics of Finnish starlings in 1951-2005 : from monitoring to population modelling

Farmland bird species have been declining in Europe. Many declines have coincided with general intensification of farming practices. In Finland, replacement of mixed farming, including rotational pastures, with specialized cultivation has been one of the most drastic changes from the 1960s to the 1990s. This kind of habitat deterioration limits the persistence of populations, as has been previously indicated from local populations. Integrated population monitoring, which gathers species-specific information of population size and demography, can be used to assess the response of a population to environment changes also at a large spatial scale. I targeted my analysis at the Finnish starling (Sturnus vulgaris). Starlings are common breeders in farmland habitats, but severe declines of local populations have been reported from Finland in the 1970s and 1980s and later from other parts of Europe. Habitat deterioration (replacement of pasture and grassland habitats with specialized cultivation areas) limits reproductive success of the species. I analysed regional population data in order to exemplify the importance of agricultural change to bird population dynamics. I used nestling ringing and nest-card data from 1951 to 2005 in order to quantify population trends and per capita reproductive success within several geographical regions (south/north and west/east aspects). I used matrix modelling, acknowledging age-specific survival and fecundity parameters and density-dependence, to model population dynamics. Finnish starlings declined by 80% from the end of the 1960s up to the end of the 1980s. The observed patterns and the model indicated that the population decline was due to the decline of the carrying capacity of farmland habitats. The decline was most severe in north Finland where populations largely become extinct. However, habitat deterioration was most severe in the southern breeding areas. The deteriorations in habitat quality decreased reproduction, which finally caused the decline. I suggest that poorly-productive northern populations have been partly maintained by immigration from the highly-productive southern populations. As the southern populations declined, ceasing emigration caused the population extinction in north. This phenomenon was explained with source sink population dynamics, which I structured and verified on the basis of a spatially explicit simulation model. I found that southern Finnish starling population exhibits ten-year cyclic regularity, a phenomenon that can be explained with delayed density-dependence in reproduction.